Oecologia (2002) 131:20–26
DOI 10.1007/s00442-001-0846-8
Abstract Many arthropods produce clusters of eggs, but
an unambiguous explanation for the evolution of egg
clustering is still lacking. We test several hypotheses for
the production of egg clusters by the predatory mite
Iphiseius degenerans. This predator feeds on pollen,
thrips larvae and nectar in flowers, but oviposits in clus-
ters in tufts of leaf hairs (acarodomatia), where eggs run
a lower risk of being killed by thrips, the prey of this
predatory mite. The observed clustering is not caused by
a shortage of oviposition sites; females preferably ovi-
posit in a domatium containing eggs rather than in an
empty domatium. To explain this preference, we first ex-
amined the effect of egg clusters on the risk of cannibal-
ism. We found that eggs are invulnerable to cannibalism,
whereas larvae emerging from single eggs or from clus-
ters were equally vulnerable. Subsequently, we consid-
ered the killing of eggs resulting from counter-attacks by
prey, i.e. the western flower thrips. We found no indica-
tion that a cluster of eggs protects eggs from predation
by thrips. However, when eggs were clustered in a dom-
atium rather than scattered over domatia, the proportion
of eggs killed by thrips was lower. Hence, oviposition in
clusters has no effect on its own and oviposition in
domatia reduces predation risk by thrips, but oviposition
in clusters in domatia leads to a synergistic effect on the
survival of predator eggs. This synergism probably arises
because eggs in clusters within tufts of leaf hairs are
more difficult for thrips to reach. These experiments
highlight a novel explanation of egg clustering, i.e. adap-
tation to counter-attacking prey. Moreover, they show
that plant domatia protect predator eggs from predation.
Keywords Domatia · Egg clustering · Predation ·
Counter-attacking prey · Cannibalism
Introduction
Oviposition in clusters is a common phenomenon in var-
ious insect taxa (Parker 1965; Weseloh 1972; Stamp
1980; Ito et al. 1982; Chew and Robbins 1984; Damman
and Cappuccino 1991; Stewart et al. 1991; McCauley
1992; Agarwala and Dixon 1993; le Masurier 1994;
Codella and Raffa 1995; Hunter 1995; Clark and Faeth
1997, 1998). Several terms have been used to describe
this form of oviposition, such as egg clumping, egg pool-
ing, aggregation of eggs or batches of eggs, but defini-
tions regarding these terms are absent or ambiguous. In
this paper, an egg cluster is defined as a group of eggs,
produced by one or more females, in which each egg
touches the rest of the group through contact with at
least one other egg.
There are three major groups of explanations for the
evolution of egg clustering. Some of these explanations
are poorly supported by published data, while others are
likely to be the consequence rather than the cause of
such oviposition behaviour (Courtney 1984). The first
group centres around the direct effects on the ovipositing
female; by reducing search for oviposition sites and lay-
ing many eggs together, the female saves energy, incurs
a lower mortality risk during searching (by predation or
adverse weather conditions), and/or reduces the risk of
failing to find another oviposition site (Stamp 1980).
These explanations implicitly assume that moving from
one oviposition site to another has associated costs and
that selection therefore acts to minimise such movement.
The second group of explanations focuses on the ben-
efits of clustering through effects on fitness of the larvae
that hatch from the eggs. Larval survivorship and perfor-
mance are thought to be enhanced by aggregation be-
cause of predator saturation (Lawrence 1990), communal
defence against predators and parasitoids (Labine 1968;
Tostowaryk 1972; Stamp 1980; Chew and Robins 1984),
an enhanced effect of warning coloration (Labine 1968;
Stamp 1980), increased efficiency of thermo-regulation
(Seymour 1974; Porter 1982), or increased efficiency of
resource use (Young and Moffett 1979; Tsubaki and
F. Faraji · A. Janssen (
✉
) · M.W. Sabelis
Section Population Biology,
Institute for Biodiversity and Ecosystem Dynamics,
University of Amsterdam, P.O. Box 94084, 1090 GB, Amsterdam,
The Netherlands
e-mail: janssen@science.uva.nl
Fax: +31-20-5257754
POPULATION ECOLOGY
Farid Faraji · Arne Janssen · Maurice W. Sabelis
The benefits of clustering eggs:
the role of egg predation and larval cannibalism in a predatory mite
Received: 20 August 2001 / Accepted: 22 October 2001 / Published online: 30 November 2001
© Springer-Verlag 2001